A CT-based Survey of Supramedullary Diverticula in Extant Birds

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1 A CT-based Survey of Supramedullary Diverticula in Extant Birds Dr. Jessie Atterholt 1,2 & Dr. Mathew Wedel 1 1Western University of Health Sciences, Pomona, CA, USA 2Raymond M. Alf Museum of Paleontology, Claremont, CA, USA

2 (Müller, 1908) lungs & trachea air sacs & diverticula

3 cervical air sacs lungs abdominal air sacs (Müller, 1908)

4 diverticula supervertebralia (Müller, 1908) canalis intertransversarius (Müller, 1908)

5 Supramedullary Airways: diverticula branching from lungs & air sacs inside vertebral canal above spinal cord

6 supramedullary airways spinal cord vertebral canal

7 supramedullary airways spinal cord vertebral canal

8 supramedullary airways spinal cord vertebral canal

9

10 Research on Supramedullary Airways Müller (1908) pigeon Cover (1953) turkey Bezuidenhout et al. (1999) ostrich O Connor (2006) general description

11 Research on Supramedullary Airways Müller (1908) pigeon Cover (1953) turkey Bezuidenhout et al. (1999) ostrich O Connor (2006) general description

12 Prior Phylogenetic Sampling Strigiformes Accipitridae Cathartidae Charadriiformes Rallidae Podicipediformes Columbiformes Gaviiformes Pteroclidiformes Musophagiformes Cuculiformes Falconiformes Coraciiformes Passeriformes Piciformes Trogoniformes Coliiformes Procellariiformes Caprimulgiformes Apodiformes Psittaciformes Pelecaniformes Gruiformes Struthioniformes Anseriformes Galliiformes 2 taxa Combined analysis tree (Atterholt & Matzke, 2011) 2 major clades

13 Broad Phylogenetic Sampling Strigiformes Accipitridae Cathartidae Charadriiformes Rallidae Podicipediformes Columbiformes Gaviiformes Pteroclidiformes Musophagiformes Cuculiformes Falconiformes Coraciiformes Passeriformes Piciformes Trogoniformes Coliiformes Procellariiformes Caprimulgiformes Apodiformes Psittaciformes Pelecaniformes Gruiformes Struthioniformes Anseriformes Galliiformes 26 genera Combined analysis tree (Atterholt & Matzke, 2011) 18 major clades

14 Supramedullary Airways Present Anas platyrhynchos (Anseriformes) Musophaga violacea (Musophagiformes) Zenaida macroura (Columbiformes) Meleagris gallopavo (Galliformes) Puffinus griseus (Procellariiformes) Bubo virginianus (Strigiformes) Struthio camelus (Struthioniformes) Calypte anna (Apodiformes) Falco sparverius (Falconiformes) Buteo jamaicensis Cathartes aura (Accipitriformes) Uria aalge Larus occidentalis (Charadriiformes) Phalacrocorax penicillatus Pelecanus occidentalis (Pelicaniformes) Nycticorax nycticorax Pyrrhura molinae Eclectus roratus (Psittaciformes) Melozone crisallis Aphelocoma californica Psaltriparus minimus Turdus migratorius (Passeriformes)

15 Supramedullary Airways Absent Gavia immer (Gaviiformes) Gavia pacifica (Gaviiformes) Gavia adamsii (Gaviiformes) Aechmorphus occidentalis (Podicipediformes) Dryobates nuttallii (Piciformes) Melanerpes formicivorus (Piciformes)

16 Supramedullary Airways Absent Aechmorphus occidentalis (Podicipediformes) Gavia pacifica (Gaviiformes) diving taxa

17 Supramedullary Airways Absent Dryobates nuttallii (Piciformes) Melanerpes formicivorus (Piciformes)???

18 Diving Taxa Aechmorphus occidentalis (Podicipediformes) Phalacrocorax penicillatus (Suliformes) Gavia pacifica (Gaviiformes) Supramedullary Airways Absent Uria aalge (Charadriiformes) Supramedullary Airways Present

19 Presence and Absence of SMAs Strigiformes Accipitridae Cathartidae Charadriiformes Rallidae Podicipediformes Columbiformes Gaviiformes Pteroclidiformes Musophagiformes Cuculiformes Falconiformes Coraciiformes Passeriformes Piciformes Trogoniformes Coliiformes Procellariiformes Caprimulgiformes Apodiformes Psittaciformes Pelecaniformes Gruiformes Struthioniformes Anseriformes Galliiformes Combined analysis tree (Atterholt & Matzke, 2011) Phylogenetic Signal 26 taxa 18 major clades

20 Morphology of Supramedullary Airways anterior dorsal view posterior anterior view spinal cord

21 Morphology of Supramedullary Airways canalis intertransversarius only (no SMAs)

22 Morphology of Supramedullary Airways A. supramedullary airways at intervertebral joints

23 Morphology of Supramedullary Airways B. supramedullary airways discontinuous through vertebral canals

24 Morphology of Supramedullary Airways C. supramedullary airways continuous through vertebral canals

25 Morphology of Supramedullary Airways D. supramedullary airways merge with diverticula supervertebralia

26 Morphology of Supramedullary Airways A B C D variation across vertebral column is universal

27 Morphology of Supramedullary Airways ostrich 2 dorsal tubes

28 Morphology of Supramedullary Airways bushtit 2 lateral tubes

29 Morphology of Supramedullary Airways ostrich 3 dorsal tubes

30 Morphology of Supramedullary Airways pelican spinal cord jacketed by SMAs

31 Variation with Body Size ostrich pelican

32 Variation with Body Size? bushtit turkey vulture

33 Variation Across Vertebral Regions cervical vertebrae present: 22 genera absent: 0 genera red-tailed hawk

34 Variation Across Vertebral Regions dorsal vertebrae present: 13 genera absent: 9 genera green-cheeked conure

35 Variation Across Vertebral Regions synsacrum *present: 5 genera absent: 17 genera great-horned owl *in posterior sacrals only

36 Variation Across Vertebral Regions caudal vertebrae present: 3 genera absent: 19 genera pelican

37 Conclusions: Variation in Supramedullary Airways 1. phylogenetic position 2. body size 3. vertebral region Strong phylogenetic signal Gavia immer (Gaviiformes) Gavia pacifica (Gaviiformes) Gavia adamsii (Gaviiformes) absent Dryobates nuttallii Melanerpes formicivorus (Piciformes) (Piciformes) Pyrrhura molinae Eclectus roratus (Psittaciformes) Buteo jamaicensis Cathartes aura (Accipitriformes) Uria aalge Larus occidentalis (Charadriiformes) Phalacrocorax penicillatus Pelecanus occidentalis (Pelicaniformes) Nycticorax nycticorax Melozone crisallis Aphelocoma californica Psaltriparus minimus Turdus migratorius (Passeriformes) present

38 Conclusions: Variation in Supramedullary Airways 1. phylogenetic position 2. body size? 3. vertebral region bushtit turkey vulture Future studies: quantify volume of SMAs and spinal cord

39 Conclusions: Variation in Supramedullary Airways 1. phylogenetic position 2. (body size) 3. vertebral region 100 % genera present cervical dorsal sacral caudal vertebral region Function influences distribution of SMAs along vertebral column? more movement > large SMAs restricted movement > small (absent) SMAs

40 Conclusions: Defining Supramedullary Airways diverticula branching from lungs & air sacs inside vertebral canal dorsal to spinal cord

41 Conclusions: Defining Supramedullary Airways diverticula branching from lungs & air sacs inside vertebral canal or at intervertebral joints dorsal to spinal cord

42 Conclusions: Defining Supramedullary Airways diverticula branching from lungs & air sacs inside vertebral canal or at intervertebral joints dorsal to spinal cord or otherwise in contact

43 Conclusions: Paramedullary Diverticula diverticula branching from lungs & air sacs inside vertebral canal or at intervertebral joints dorsal to spinal cord or otherwise in contact

44 Osteological Correlates foramina sculpting/fossae Phoebastria nigripes LACM Cervical 9 length 20mm Pelecanus erythrorhynchus LACM Cervical 13 length 40mm

45 Osteological Correlates pelican foramen in floor of vertebral canal

46 Osteological Correlates foramina sculpting/fossae (Schwarz & Fritsch, 2006: Fig. 4) Giraffatitan Alamosaursus

47 Conclusions: Paramedullary Airways in Other Archosaurs? Illustration by Brian Engh (dontmesswithdinosaurs.com)

48 Conclusions: Paramedullary Airways in Other Archosaurs? Photo: Dave Hone Pteranodon (LACM 51760) foramina in floor of vertebral canal

49 Thank you! Illustration by Brian Engh (dontmesswithdinosaurs.com) Questions?

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